Scorpaenodes guamensis ( Quoy & Gaimard 1824 )

Motomura, Hiroyuki, Causse, Romain & Struthers, Carl D., 2016, Redescription of the Indo-Pacific scorpionfish Scorpaenodes guamensis (Quoy & Gaimard 1824) (Scorpaenidae), a senior synonym of seven nominal species, Zootaxa 4067 (3), pp. 345-360 : 346-356

Scorpaenodes guamensis ( Quoy & Gaimard 1824)

[English name: Guam Scorpionfish]

Figures 1 A–G; Tables 1–3

Scorpaena guamensis Quoy & Gaimard 1824: 326 (type locality: Guam, Mariana Islands).

Scorpaena rubropunctata Cuvier in Cuvier & Valenciennes 1829: 324 (type locality: Red Sea).

Sebastes minutus Cuvier in Cuvier & Valenciennes 1829: 348 (original locality: Guam, Moluccas, Waigiou, Timor, Tahiti, Bora Bora; type locality: Guam, based on lectotype).

Scorpaena chilioprista Rüppell 1838: 107 , pl. 27, fig. 3 (type locality: Massawa, Eritrea, Red Sea).

Scorpaena polylepis Bleeker 1851: 173 (type locality: western Sumatra, Indonesia).

Centropogon echinatus Macleay 1881: 436 (type locality: Endeavour River, Queensland, Australia).

Scorpaena erinacea Garman 1903: 231 , pl. 1, fig. 2 (type locality: Suva Reef, Viti Levu, Fiji).

Scorpaenopsis quiescens Seale 1906: 80 , fig. 2 (type locality: Tahiti, Society Islands).

Material examined. Lectotype. MNHN 6667, 44.0 mm SL, Guam, Mariana Islands, J. R. C. Quoy & J. P. Gaimard. [Designated herein].

Paralectotype. MNHN 6667, 29.5 mm SL, same data as lectotype.

Other type material. 23 specimens, 29.5–79.9 mm SL. Sebastes minutus Cuvier 1829 : MNHN 6666, paralectotype, 57.2 mm SL, Molucca Islands, Indonesia, F. Péron and C. A. Lesueur, 1804; MNHN 6667, lectotype (herein designated; also lectotype of Scorpaena guamensis Quoy & Gaimard 1824 ), 44.0 mm SL; MNHN 6667, paralectotype (also paralectotype of S. guamensis ), 29.5 mm SL; MNHN 6668, 2 paralectotypes, 53.8–60.8 mm SL, Tahiti, Society Islands, R. P. Lesson & P. Garnot; MNHN 6669, 2 paralectotypes, 44.1–51.3 mm SL, Bora Bora, Society Islands, R. P. Lesson & P. Garnot; MNHN 6670, paralectotype, 79.9 mm SL, Waigiou, Indonesia, J.

R. C. Quoy & J. P. Gaimard; RMNH 657, paralectotype, 51.5 mm SL, Bora Bora, Society Islands; SMF 1473, paralectotype, 58.8 mm SL, Timor, Indonesia (other paralectotype of S. minutus , 48.2 mm SL, registered as SMF 1473 identified as Scorpaenodes varipinnis ). Scorpaena rubropunctata Cuvier 1829 : ZMB 765, 4 paralectotypes, 37.1–43.4 mm SL, Red Sea; ZMB 766, lectotype (designated herein), 32.6 mm SL, Red Sea; ZMB 766, paralectotype, 33.6 mm SL, Red Sea. Scorpaena chilioprista Rüppell 1838 : SMF 476, holotype, 50.2 mm SL, Massawa, Eritrea, Red Sea, E. Rüppell. Scorpaena polylepis Bleeker 1851 : RMNH 5855, lectotype (designated herein), 32.8 mm SL, western Sumatra, Indonesia, P. Bleeker; RMNH 35642, paralectotype, 34.9 mm SL, same data as RMNH 5855; SMNS 10633, paralectotype, 41.2 mm SL, western Sumatra, Indonesia, P. Bleeker. Centropogon echinatus Macleay 1881 : AMS I. 16353-001, holotype, 69.8 mm SL, Endeavour River, Queensland, Australia. Scorpaena erinacea Garman 1903 : MCZ 28313, holotype, 57.5 mm SL, Suva Reef, Viti Levu, Fiji. Scorpaenopsis quiescens Seale 1906 : BPBM 1352, holotype, 71.7 mm SL, Tahiti, Society Islands, A. Seale, Feb. 1901.

Non-type material. 112 specimens, 11.6–84.6 mm SL. RED SEA: MNHN 1952-272, 38.7 mm SL, 21°30′00″N, 37°34′59″E, Abulat, RV Calypso ; SMF 15634, 50.1 mm SL, Massawa, Eritrea, Red Sea, E. Rüppell (previously treated as a possible paratype of Scorpaena chilioprista Rüppell, 1838 ). GULF OF TADJOURA: MNHN 1977-742, 38.2 mm SL, Maskali Islands, off Djibouti, 11°45′00″N, 43°10′01″E, 20 m, J. E. Randall & A. L. Maugé, May 1977. MASCARENE ISLANDS: USNM 349955, 72.0 mm SL, Trou Aux Biches, Mauritius, 4–5 m, A. Gill et al., 9 May 1995. JAPAN: KAUM –I. 11581, 69.8 mm SL, Harutahama Beach, Ambo, Yaku-shima, Osumi Islands, Kagoshima, 30°18′02″N, 130°39′17″E, 0.5–1.5 m, M. Meguro, 18 Aug. 2008; KAUM –I. 17391, 75.8 mm SL, KAUM –I. 17400, 79.3 mm SL, tide-pool at Tsuchihama, Kasari, Amami-oshima, Amami Islands, Kagoshima, 28°24′34″N, 129°40′31″E, 0.5–1.0 m, M. Meguro & T. Yoshida, 26 Mar. 2009; KAUM –I. 51550, 64.5 mm SL, off Maehama beach, south coast of Yoron-jima, Amami Islands, Kagoshima, 27°01′13″N, 128°26′26″E, 2 m, KAUM fish team, 29 Oct. 2012; KAUM –I. 53825, 70.0 mm SL, Nagura Bay, Ishigaki-jima, Yaeyama Islands, Okinawa, 24°24′51″N, 124°07′31″E, 1 m, T. Yoshida & H. Nishiyama, 5 Apr. 2012; KPM-NI 3059, 31.3 mm SL, KPM-NI 3099, 22.0 mm SL, KPM-NI 3114, 14.6 mm SL, KPM-NI 3135, 40.3 mm SL, KPM-NI 3139, 33.8 mm SL, KPM-NI 3144, 26.2 mm SL, KPM-NI 3167, 2, 28.5–35.9 mm SL, Hachijo-jima island, Izu Islands, H. Senou et al., 24 Nov. 1996; KPM-NI 4725, 61.3 mm SL, Ie-jima island, Okinawa Islands, Ryukyu Islands, 15 June 1998; KPM-NI 7346, 11.9 mm SL, Jogashima, Miura, Kanagawa, K. Yamada, 16 Sept. 1999; KPM-NI 8470, 23.3 mm SL, Ie-jima island, Okinawa Islands, Ryukyu Islands, T. Nomura, 18 Sept. 2001; KPM-NI 10669, 48.5 mm SL, KPM-NI 10670, 47.1 mm SL, Ie-jima island, Okinawa Islands, Ryukyu Islands, T. Nomura & M. Takahashi, 28 July 2002; KPM-NI 13464, 19.3 mm SL, Amashiru, Zamami Island, Kerama Islands, Ryukyu Islands, 5 m, K. Uchino, 5 June 2004; KPM-NI 20164, 55.4 mm SL, KPM-NI 20219, 54.9 mm SL, Oganeku, Yoron-jima island, Amami Islands, A. Kishimoto, 21 Aug. 2007. TAIWAN: KAUM –I. 39263, 62.4 mm SL, Wan Li Tong, Kenting, Pingdong, 21°59′52″N, 120°42′03″E, 0.5 m. MARIANA ISLANDS: USNM 139894, 46.5 mm SL, Guam, F. Craighead & J. Craighead, Nov. 1945. MARSHALL ISLANDS: MNHN 1976-339, 5, 51.7–66.6 mm SL, Enewetak, 11°30′00″N, 162°30′00″E, M.-L. Bauchot & J. E. Randall, Apr. 1976. CAROLINE ISLANDS: MCZ 29468, 44.5 mm SL, 05°25′N, 163°00′E, RV Albatross, A. Agassiz, 9 Feb. 1900; USNM 224494, 1 of 2, 53.5 mm SL, Nanmatol, Ponape, 05°51′N, 158°20′E, 2 m, V. Springer et al., 3 Sept. 1980. PHILIPPINES: USNM 372635, 48.9 mm SL, Sombrero Island, Batang Islands, 13°42′N, 120°49′E, 6 m, 24 Apr. 1980. MALAYSIA: KPM-NI 1991, 43.6 mm SL, Mabul Island, Sabah, Borneo. INDONESIA: AMS B. 8267, 69.9 mm SL, Ambon, Indonesia, F. Day, 1883 (previously treated as a syntype of Scorpaena polylepis Bleeker 1851 ); RMNH 35643, 8, 22.5–81.7 mm SL, western Sumatra, Indonesia, P. Bleeker (previously treated as possible syntypes of Scorpaena polylepis Bleeker 1851 ); SMNS 10633, 62.9 mm SL, western Sumatra, Indonesia, P. Bleeker (previously treated as a possible syntype of Scorpaena polylepis Bleeker 1851 ); USNM 210946, 1 of 2, 55.9 mm SL, Tandjung Tala, Nusa Laut, Moluccas, 16 Jan. 1973; USNM 422288, 58.6 mm SL, Ambon Bay, Ambon, Moluccas, 03°39′00″S, 128°11′30″E, 2 m, V. Springer & M. Gomon, 17 Mar. 1974. PAPUA NEW GUINEA: AMS I. 17504-030, 48.4 mm SL, Japanese submarine base, off Cape Tawui, 04°07′S, 152°14′E, 1–5 m, G. R. Allen, 9 Aug. 1973; USNM 122985, 1 of 3, 53.3 mm SL, Bougainville Island, Aug. 1944; USNM 374919, 1 of 2, 53.1 mm SL, Meman Island, Ninigo Islands, 01°12′40″S, 144°16′59″E, 1.2 m, V. Springer et al., 12 Nov. 2004. SOLOMON ISLANDS: USNM 422296, 38.8 mm SL, Munde Reef, New Georgia, W. Chapman, 12 May 1944. NEW HEBRIDES: USNM. 285930, 50.7 mm SL, Santo, A. Power, 1986. AUSTRALIA: MCZ 36909, 5, 28.8–68.9 mm SL, off Cairns, Queensland, O. Barton; MCZ 90605, 75.7 mm SL, Rib Reef, off Townsville, Queensland, 0–10 m, D.

Williams, 1984. NEW CALEDONIA: MNHN 1980-150, 3, 44.5–48.9 mm SL, Mare, Loyalty Islands, Nov. 1970; MNHN 1980-390, 49.8 mm SL, 21°30′00″S, 165°25′01″E, P. Fourmanoir, Dec. 1970; MNHN 1980-809, 36.2 mm SL, 21°45′S, 164°15′E, 12 m, M.-L. Bauchot & A. L. Maugé, Jan. 1979; MNHN 2005-2628, 4, 40.6–47.9 mm SL, 22°26′20″S, 166°24′14″E, M. Kulbicki, 2 Sept. 1985; NMNZ P.027308, 53.4 mm SL, NMNZ P.027373, 2, 56.0– 59.1 mm SL, Tabu Reef, Passe de Boulari, 22°28′59″S, 166°26′17″E, 10–18 m, C. Roberts, 16 Oct. 1991; NMNZ P.027358, 59.2 mm SL, off Laregnere Reef, southwest of Nouméa, 22°19′59″S, 166°17′30″E, 12–17 m, C. Roberts, 17 Oct. 1991; NMNZ P.029459, 79.1 mm SL, Seche Croissant Reef, west-southwest of Nouméa, 22°19′00″S, 166°20′59″E, 8–10 m, C. Roberts & C. Paulin, 29 Oct. 1992; USNM 319727, 62.1 mm SL, off Nouméa, W. Chapman et al., 13 Apr. 1944. FIJI: NMNZ P.014611, 69.1 mm SL, Makuluvu, 17°30′00″S, 179°00′00″E, L. Simpson, Jan. 1958; USNM 244009, 51.8 mm SL, off Malamala Island, 17°44′S, 177°17′E, 0–7.6 m, V. Springer et al., 25 May 1982. SAMOA: MCZ 30246, 3, 54.0– 54.7 mm SL, Apia; MCZ 30273, 46.9 mm SL, Apia, 1902. TONGA: USNM 338543, 57.1 mmm SL, Ovaka Island, 18°44′31′S, 174°06′37″W, 24–29 m, J. Williams et al., 17 Nov. 1993. AMERICAN SAMOA: USNM 115712, 64.4 mm SL, Pago Pago Bay, Tutuila Island, L. Schultz, 2 June 1939. NIUE: NMNZ P.014592, 84.6 mm SL, 19°1′59″S, 169°55′14″W, W. McDowall, Aug. 1962. TUVALU: NMNZ P.044942, 52.9 mm SL, NMNZ P.044943, 48.8 mm SL, NMNZ P.044944, 30.4 mm SL, NMNZ P.044945, 29.2 mm SL, Funafuti Lagoon, Funafuti, 08°35′58″S, 179°05′27″E, 2–6 m, C. Struthers et al., 10 Sept. 2008; NMNZ P.045008, 39.3 mm SL, NMNZ P.045009, 60.2 mm SL, NMNZ P.045010, 63.3 mm SL, NMNZ P.045011, 63.2 mm SL, NMNZ P.045012, 63.0 mm SL, NMNZ P.045013, 66.4 mm SL, Funafuti Lagoon, Funafuti, 08°32′38″S, 179°09′49″E, 6–9 m, C. Struthers et al., 12 Sept. 2008. FRENCH POLYNESIA: AMS I. 33667-016, 66.0 mm SL, Mataiva Atoll, Tuamotu Archipelago, 14°49′S, 148°34′W, J. Bell, Sept. 1983; MNHN 1984-282, 16, 11.6–67.9 mm SL, Papetoai, Moorea, Society Islands, 17°28′59″S, 149°52′01″W, 2 m, R. Galzin, Aug. 1982; MNHN 2008-388, 22.8 mm SL, Moorea, Society Islands, 1 m, 11 Mar. 2006; MNHN 2008-633, 66.9 mm SL, MNHN 2008-634, 49.5 mm SL, MNHN 2008-635, 39.3 mm SL, Paroa, Moorea, Society Islands, 1 m, 15 Mar. 2006; USNM 379833, 61.7 mm SL, Anarua Bay, Rapa, 27°36′18″S, 144°22′48″W, 2 m, J. Williams et al., 7 Nov. 2002.

Diagnosis. A species of Scorpaenodes with the following combination of characters: 17–20 (mode 18) pectoral-fin rays, some rays branched in specimens over ca. 40 mm SL; 7–9 (8) dorsal-fin soft rays; 40–46 (43) scale rows in longitudinal series; no scales on underside of lower jaw; 11–15 (13) gill rakers; nasal spine present; lateral lacrimal spine usually present; 2 suborbital spines, first spine below eye, second spine at end of suborbital ridge; interorbital and coronal spines absent; large dark blotch on opercle; no dark blotch on subopercle or spinous portion of dorsal fin.

Description. Data for the lectotype are presented first, followed by data for other materials (if different) in parentheses; descriptions of caudal-fin rays and swimbladder are based on non-type specimens. Dorsal fin with 13 spines (rarely 14 spines); fifth (or sixth) spine longest, its length subequal to pelvic-fin spine length; spines six to twelve becoming progressively shorter posteriorly; length of twelfth spine 1.4 (1.0–1.5, mean 1.2) in last spine; membrane of spinous portion of dorsal fin moderately notched. Dorsal fin with 8 soft rays (rarely 7, if 14 dorsal-fin spines; rarely 9 rays, only 7% of specimens examined in this study); soft rays damaged in lectotype (all rays branched, divided into 2–4 branches; second soft ray longest, its length slightly shorter than longest anal-fin soft ray length); posterior branch of last ray not joined by membrane to caudal peduncle (joined by membrane to caudal peduncle for approximately one-third to one-fourth its length). Anal fin with 3 spines; second spine longest and widest, its length shorter than pelvic-fin spine length; first spine 2.5 (1.8–2.7, mean 2.3) in second spine, 1.9 (1.5– 2.2, mean 1.9) in third spine. Anal fin with 5 soft rays; all rays branched, divided into 2 (2–4) branches; first ray longest; posterior branch of last ray not joined by membrane to caudal peduncle (joined by membrane to caudal peduncle for approximately one-third its length). Pectoral fin with 18 rays on each side of body (17–20); upper 2 rays and lower 11 rays unbranched, remaining middle rays branched (all rays unbranched in small specimens usually less than about 40 mm SL; number of branched rays increasing up to 8 rays with growth, but uppermost ray always unbranched); tenth (rarely ninth) ray longest, its length subequal to body depth; lower rays not covered with thick skin; posterior margin of fin not bilobed. Pelvic fin with 1 spine and 5 soft rays; last ray unbranched (branched in larger specimens), other rays branched, divided into 2 (2–8) branches; second ray longest, its length longer than longest anal-fin soft ray length; last soft ray joined by membrane to abdomen for more than half its length. Four (rarely 5) dorsal series of caudal-procurrent rays, 4 ventral series; 2 (rarely 1) dorsal series of segmented, unbranched caudal-fin rays, 2 (rarely 3) ventral series; 6 (rarely 7) dorsal series of branched caudal-fin rays, 6 (rarely 4 or 5) ventral series. Posterior margin of caudal fin slightly rounded. Formula for configuration of anterior neural spines and anterior dorsal pterygiophores ///2+1/1/1/1/1/1/1/1/1/1/1. Caudal-peduncle depth 1.6 (1.1–1.7, mean 1.4) in caudal-peduncle length.

Scorpaenodes guamensis

S. guamensis and S. minutus S. rubropunctata S. minutus ......continued on the next page Longest dorsal-fin soft ray length - - - - 16.9 1rakers on upper limb + on ceratohyal + on hypobranchial = total; 2at vertical midline of eye; 3at posterior end of preocular spine base; 4counts and measurements based on 5 and 2 specimens, respectively (see text).

Pored lateral-line scales 23 (22–25, mode 23). Scale rows above lateral line 5 (4–7, mode 5), below lateral line 10 (10–14, mode 12). Scale rows between sixth dorsal-fin spine base and lateral line 5 (4–6, mode 5), between last dorsal-fin spine base and lateral line 4 (4–6, mode 5). Predorsal scale rows 7 (5–8, mode 6). Gill rakers on upper limb 6 (4–7, mode 6), lower limb 7 (6–9, mode 7), total gill rakers 13 (11–15, mode 13); no raker on hypobranchial (rarely 1 or 2 rudimentary rakers); length of longest raker on first gill arch subequal to length of gill filaments around angle of gill arch; no slit behind lower fourth gill arch. Branchiostegal rays 7. Swimbladder absent.

Body strongly compressed anteriorly, progressively more compressed posteriorly. Nape and anterior body not strongly arched. Body depth relatively shallow; less than head length. Few small papillae scattered on upper margin of eye membrane. Short, broad tentacle on posterior edge of low membranous tube associated with anterior nostril; tentacle extending beyond anterior margin of posterior nostril when laid back. Supraocular tentacle slender, short, its length less than pupil diameter (often broad, with several short branches, and long, its length slightly greater than pupil diameter). Posterior lacrimal spine not linked posteriorly to head by skin.

Exposed ctenoid scales covering interorbital space (partly), occipital region, and area between parietal and pterotic spine bases in dorsal view; well-exposed ctenoid scales on lateral surface of head, except for snout region, lacrimal, maxilla, and subopercle. Well-exposed ctenoid scales on lateral surface of body, scales becoming cycloid on abdomen. Body scales extending onto spines, rays or membranes of fins, except pelvic fin. Exposed cycloid scales (some with weak cteni) covering pectoral-fin base and ventral surface of body, including between pelvic fins.

Cephalic sensory pores prominent; 3 circular pores just below suborbital ridge (pores often covered with thin skin); first pore located just behind lacrimal bone, second pore below second suborbital spine, and third pore below end of suborbital ridge. Anterior nostril circular, located just below nasal spine base, its diameter less than onethird of posterior nostril diameter. Posterior nostril circular, located behind nasal spine. Pair of elliptic pores on interorbital space in dorsal view (usually covered with thin skin); each pore located anterior to midline through pupil and between interorbital ridge and supraocular ridge. Several elliptic pores along preopercular margin mostly covered with thin skin. Underside of dentary with 3 sensory pores on each side in ventral view; first pore below anterior edge of lacrimal bone, second pore below and between tips of anterior and posterior lacrimal spines, third pore located on posterior margin of dentary. Pore behind symphysial knob of lower jaw on each side.

Mouth large, slightly oblique, forming angle of about 20 degrees to horizontal axis of head and body. Posterior margin of maxilla just reaching (or extending slightly beyond) vertical through posterior margin of orbit. Posterior upper edge of maxilla slightly swollen laterally; anterior part of maxilla slightly convex, central part nearly flat. Lower jaw with a symphysial knob. Width of symphysial gap separating premaxillary teeth bands subequal to width of each band. Upper jaw with a band of short, incurved, conical teeth, tips of teeth not strongly pointed. Tooth band in about 8 (up to 18) teeth at front of upper jaw, narrowing posteriorly; band in upper jaw wider than that of lower jaw. Lower jaw with a band of short, incurved, conical teeth; lengths of most teeth subequal to those of upper jaw. Seven (up to 10) rows of small teeth at front of vomer, becoming 3 (3–5) rows posteriorly, forming Vshaped patch. No palatine teeth. Underside of lower jaw smooth without ridges.

Scorpaenodes guamensis ......continued on the next page Scorpaenodes guamensis 1rakers on upper limb + on ceratohyal + on hypobranchial = total; 2at vertical midline of eye; 3at posterior end of preocular spine base.

Dorsal profile of snout elevated slightly, forming an angle of about 20 degrees to horizontal axis of head and body. Nasal spine simple, sharp, directed dorsally, its length greater than anterior nostril diameter. Ascending process of premaxilla not strongly intruding into interorbital space, its posterior margin not extending beyond line between posterior ends of preocular spine bases in dorsal view. Median interorbital ridge absent. Interorbital ridges poorly developed, separated by relatively shallow furrow, beginning posterior to nasal spines and not posteriorly joining tympanic spine bases; interorbital ridges diverging anteriorly and posteriorly in dorsal view, distance between interorbital ridges narrowest slightly anterior to vertical midline through eye. Interorbital space relatively shallow, about one-tenth of orbit diameter, extending above dorsal profile of head. Preocular spine simple, directed dorsoposteriorly; tip of spine extending beyond level of upper margin of pupil in lateral view. Supraocular spine simple, its tip reaching vertical midline through pupil in lateral view; spine shorter than postocular and tympanic spines. Postocular spines simple, slightly canted laterally. Tympanic spine simple, strongly pointed, directed dorsoposteriorly. Interorbital, coronal and pretympanic spines absent. Indistinct transverse ridge anterior to occiput slightly curved posteromedially in dorsal view. Occiput nearly flat, lacking pit. No transverse ridge in rear of occiput between bases of parietal and nuchal spines. Nuchal and parietal spines joined at base. Sphenotic with several small spines. Postorbital with several small spines (often spines absent). Pterotic spine simple, located below bases of parietal and nuchal spines. No spine on area surrounded by parietal, nuchal, pterotic and lower posttemporal spines, but short oblique ridge on area. Upper posttemporal spine absent. Lower posttemporal spine simple, its base length less than that of pterotic spine. Supracleithral spine simple, strongly pointed; spine greater than preocular, supraocular, postocular, tympanic, parietal, nuchal, pterotic, and lower posttemporal spines. Cleithral spine flattened, strongly pointed.

Lateral lacrimal spine absent, but distinct ridge present (usually spine present). Anterior lacrimal spine triangular, simple, not pointed, directed ventrally. Posterior lacrimal spine simple, pointed, directed ventroposteriorly; posterior lacrimal spine much greater than anterior spine. Two suborbital ridges with 2 spines; first spine at end of first suborbital ridge, located below and between posterior margins of pupil and orbit, second spine at end of ridge behind orbit. Space between ventral margin of eye and suborbital ridge very narrow. Suborbital pit absent. Preopercle with 3 spines (often fourth and fifth reduced spines present in smaller specimens; 5 well developed spines in small juveniles less than 20 mm SL); uppermost spine largest with supplemental preopercular spine on its base; second spine smallest (often absent), located closer to first spine than third spine; third spine with broad base. Preopercle, between uppermost preopercular spine and upper end of preopercle, smooth without serrae or spines. Upper opercular spine simple without distinct median ridge; posterior tip of spine not reaching opercular margin. Lower opercular spine simple with a distinct median ridge; posterior tip of spine extending slightly beyond opercular margin. Space between upper and lower opercular spines without ridges.

Scorpaenodes guamensis Scorpaenodes scaber ......continued on the next page 1rakers on upper limb + on ceratohyal + on hypobranchial = total; 2at vertical midline of eye; 3at posterior end of preocular spine base; 4counts based on all specimens and measurements based on specimens over 39 mm SL; 5modes for counts and means for measurements, including data for all type and non-type specimens identified as S. guamensis ( Tables 1–2 ).

Origin of first dorsal-fin spine above supracleithral spine. Posterior margin of opercular membrane extending beyond vertical through third dorsal-fin spine base. Posterior tip of pectoral fin extending beyond vertical through first anal-fin spine base. Origin of pelvic-fin spine slightly anterior to (or just below) origin of pectoral fin. Posterior tip of depressed pelvic fin not reaching (just short of, reaching, or extending slightly beyond) anus. Origin of first anal-fin spine just below origin of last dorsal-fin spine.

Color when fresh. Based on color photographs of KAUM specimens ( Fig. 2 ). Body brownish, mottled with black, brown, or reddish brown, reddish blotches; irregular whitish vertical band usually present on caudal peduncle. Opercle with large black blotch, its size greater than pupil diameter (sometimes greater than orbit diameter); no distinct blotches on subopercle. Spinous portion of dorsal fin without distinct black blotch. Distal two-thirds of soft rayed portion of dorsal fin semitransparent with scattered reddish botches. Pectoral fin yellowish with numerous reddish blotches. Pelvic fin yellowish with dark red spots. Anal fin with reddish blotches basally, and reddish spots distally. Caudal fin semitransparent with scattered reddish blotches and/or spots.

Color of preserved specimens. In lectotype and long-term preserved specimens, opercle with large pale brown blotch; pelvic fin dark brown distally; remaining parts of body and all other fins whitish. In relatively recently collected specimens, body with numerous blackish blotches and spots; distinct large black blotch on opercle; no distinct black blotch on subopercle or spinous portion of dorsal fin; no distinct band at pectoral-fin base.

Distribution. Scorpaenodes guamensis is widely distributed in the Indo-Pacific where it ranges from the east coast of Africa and the Red Sea east to the Society Islands ( Poss, 1999; this study). This species inhabits rocky and coral reefs in depths of less than 20 m.

Figures

FIGURE 1. Primary type specimens identified as (A – G) Scorpaenodes guamensis and (H) Scorpaenodes scaber. A, lectotype of Scorpaena guamensis and Sebastes minutus (MNHN 6667, 44.0 mm SL, Mariana Islands); B, lectotype of Scorpaena rubropunctata (ZMB 766, 32.6 mm SL, Red Sea); C, holotype of Scorpaena chilioprista (SMF 476, 50.2 mm SL, Red Sea); D, lectotype of Scorpaena polylepis (RMNH 5855, 32.8 mm SL, Indonesia); E, holotype of Centropogon echinatus (AMS I. 16353 - 001, 69.8 mm SL, Australia); F, holotype of Scorpaena erinacea (MCZ 28313, 57.5 mm SL, Fiji); G, holotype of Scorpaenopsis quiescens (BPBM 1352, 71.7 mm SL, Society Islands); H, syntype of Sebastes scaber (AMS B. 8450, 41.6 mm SL, Australia).

FIGURE 2. Color photograph of fresh specimen of Scorpaenodes guamensis (KAUM – I. 51550, 64.5 mm SL, off Maehama beach, south coast of Yoron-jima, Amami Islands, Kagoshima, Japan).